Artemis (protein)

Artemis is a protein that in humans

is encoded by the DCLRE1C (DNA
DCLRE1C
cross-link repair 1C) gene.[5][6] Available structures
PDB Ortholog search: PDBe (https://www.ebi.ac.uk/pdbe/search
Results.html?display=both&term=Q8K4J0%20or%20Q96SD
Contents 1%20or%20X6R9W9) RCSB (http://www.rcsb.org/pdb/searc
h/smartSubquery.do?smartSearchSubtype=UpAccessionId
Function
Query&accessionIdList=Q8K4J0,Q96SD1,X6R9W9)
Immune response
Repair of DNA breaks List of PDB id codes

Clinical significance
Interactions
References
Further reading
External links
Function
Artemis is a nuclear protein that is
involved in V(D)J recombination and
DNA repair. The protein has
endonuclease activity on 5' and 3'
overhangs and hairpins when
complexed with PRKDC.[7]
3W1B (https://www.rcsb.org/structure/3W1B), 3W1G
(https://www.rcsb.org/structure/3W1G), 4HTP (https://
www.rcsb.org/structure/4HTP)
Identifiers
Aliases DCLRE1C (https://www.genenames.org/data/gene-symb
ol-report/#!/hgnc_id/17642), A-SCID, DCLREC1C, RS-
SCID, SCIDA, SNM1C, DNA cross-link repair 1C
External OMIM: 605988 (https://omim.org/entry/605988) MGI:
IDs 2441769 (http://www.informatics.jax.org/marker/MGI:244
1769) HomoloGene: 32547 (https://www.ncbi.nlm.nih.go
v/entrez/query.fcgi?cmd=Retrieve&db=homologene&dopt
=HomoloGene&list_uids=32547) GeneCards: DCLRE1C
(https://www.genecards.org/cgi-bin/carddisp.pl?gene=DC
LRE1C)

Gene location (Human)

Immune response

Artemis plays an essential role in

V(D)J recombination, the process by
which B cell antibody genes and T
cell receptor genes are assembled
from individual V (variable), D
(diversity), and J (joining) Chr. Chromosome 10 (human)[1]
segments.[8] For example, in joining
a V segment to a D segment, the
RAG (recombination activating
gene) nuclease cuts both DNA
strands adjacent to a V segment and Band 10p13 Start 14,897,359 bp[1]
adjacent to a D segment. The End 14,954,432 bp[1]
intervening DNA between the V and
D segments is ligated to form a Gene location (Mouse)
circular DNA molecule that is lost
from the chromosome. At each of the
two remaining ends, called the
coding ends, the two strands of DNA
are joined to form a hairpin structure.
Artemis nuclease, in a complex with
the DNA-dependent protein kinase
(DNA‑PK), binds to these DNA Chr. Chromosome 2 (mouse)[2]
ends and makes a single cut near the
tip of the hairpin. The exposed 3'
termini are subject to deletion and
addition of nucleotides by a variety
of exonucleases and DNA Band 2|2 A1 Start 3,424,131 bp[2]
polymerases, before the V and D End 3,464,130 bp[2]
segments are ligated to restore the
integrity of the chromosome. The RNA expression pattern
exact site of cleavage of the hairpin
by Artemis is variable, and this
variability, combined with random
nucleotide deletion and addition,
confers extreme diversity upon the
resulting antibody and T-cell receptor
genes, thus allowing the immune
system to mount an immune response
to virtually any foreign antigen.[9] In
Artemis-deficient individuals, V(D)J
recombination is blocked because the
hairpin ends cannot be opened, and
so no mature B or T cells are
produced, a condition known as
severe combined immune deficiency
(SCID). Artemis was first identified
as the gene defective in a subset of
SCID patients that were unusually
sensitive to radiation.

More reference expression data (http://biogps.org/gene/64421/)

Repair of DNA breaks
Cells deficient in Artemis are more
Molecular
sensitive than normal cells to
function
X‑rays[5] and to chemical agents that
induce double-strand breaks
(DSBs), [10] and they show a higher
incidence of chromosome breaks
following irradiation.[11] Direct
measurement of DSBs by pulsed-
field electrophoresis indicates that in
Artemis-deficient cells 75-90% of
DSBs are repaired rapidly, just as in
normal cells. However, the remaining
10-20% of DSBs that are repaired
more slowly (2-24 hr) in normal
Gene ontology
• 5'-3' exonuclease activity (http://amigo.geneontolog
y.org/amigo/term/GO:0008409)
• 5'-3' exodeoxyribonuclease activity (http://amigo.gen
eontology.org/amigo/term/GO:0035312)
• damaged DNA binding (http://amigo.geneontology.or
g/amigo/term/GO:0003684)
• single-stranded DNA endodeoxyribonuclease activity
(http://amigo.geneontology.org/amigo/term/GO:00000
14)
• GO:0001948 protein binding (http://amigo.geneontol
ogy.org/amigo/term/GO:0005515,)
• nuclease activity (http://amigo.geneontology.org/ami
cells, are not repaired at all in
Artemis-deficient cells.[12] Repair of
these presumably difficult-to-rejoin
breaks also requires several other
proteins, including the
Mre11/Rad50/NBS1 complex, the
ataxia telangiectasia mutated ATM
kinase, and 53BP1. Because Artemis
can remove damaged ends from
DNA,[10] it has been proposed that
these DSBs are those whose
damaged ends require trimming by
Artemis. However, evidence that
both ATM and Artemis are
specifically required for repair of
DSBs in heterochromatin,[13][14] has
called this interpretation into
question.
Artemis functions in the repair of
DNA double-strand breaks that arise
by induced oxidative reactions, or by
endogenous reactions.[15] Such
DNA repair occurs in
heterochromatin as well as in
euchromatin.
Clinical significance
Mutations in this gene cause
Athabascan-type severe combined
immunodeficiency (SCIDA).[16]
Interactions
DCLRE1C has been shown to
interact with DNA-PKcs.[17]
References
1. GRCh38: Ensembl release
89: ENSG00000152457 (htt
p://May2017.archive.ensemb
l.org/Homo_sapiens/Gene/Su
mmary?db=core;g=ENSG000
00152457) - Ensembl, May
2017
go/term/GO:0004518)
• endonuclease activity (http://amigo.geneontology.or
g/amigo/term/GO:0004519)
• exonuclease activity (http://amigo.geneontology.org/
amigo/term/GO:0004527)
• hydrolase activity (http://amigo.geneontology.org/ami
go/term/GO:0016787)
Cellular • nonhomologous end joining complex (http://amigo.g
component eneontology.org/amigo/term/GO:0070419)
• nuclear chromosome, telomeric region (http://amigo.
geneontology.org/amigo/term/GO:0000784)
• cell nucleus (http://amigo.geneontology.org/amigo/te
rm/GO:0005634)
• nucleoplasm (http://amigo.geneontology.org/amigo/t
erm/GO:0005654)
• Golgi apparatus (http://amigo.geneontology.org/amig
o/term/GO:0005794)
Biological • response to ionizing radiation (http://amigo.geneonto
process logy.org/amigo/term/GO:0010212)
• DNA recombination (http://amigo.geneontology.org/a
migo/term/GO:0006310)
• adaptive immune response (http://amigo.geneontolo
gy.org/amigo/term/GO:0002250)
• interstrand cross-link repair (http://amigo.geneontolo
gy.org/amigo/term/GO:0036297)
• protection from non-homologous end joining at
telomere (http://amigo.geneontology.org/amigo/term/
GO:0031848)
• immune system process (http://amigo.geneontology.
org/amigo/term/GO:0002376)
• chromosome organization (http://amigo.geneontolog
y.org/amigo/term/GO:0051276)
• cellular response to DNA damage stimulus (http://am
igo.geneontology.org/amigo/term/GO:0006974)
• V(D)J recombination (http://amigo.geneontology.org/
amigo/term/GO:0033151)
• B cell differentiation (http://amigo.geneontology.org/a
migo/term/GO:0030183)
• double-strand break repair via nonhomologous end
joining (http://amigo.geneontology.org/amigo/term/G
O:0006303)
• telomere maintenance (http://amigo.geneontology.or
g/amigo/term/GO:0000723)
• nucleic acid phosphodiester bond hydrolysis (http://a
migo.geneontology.org/amigo/term/GO:0090305)
• double-strand break repair (http://amigo.geneontolo
2. GRCm38: Ensembl release
89: ENSMUSG00000026648
(http://May2017.archive.ense
mbl.org/Mus_musculus/Gene/
Summary?db=core;g=ENSM
USG00000026648) -
Ensembl, May 2017
3. "Human PubMed Reference:"
(https://www.ncbi.nlm.nih.gov/
sites/entrez?db=gene&cmd=
Link&LinkName=gene_pubm
ed&from_uid=64421).
National Center for
Biotechnology Information,
U.S. National Library of
Medicine.
4. "Mouse PubMed Reference:"
(https://www.ncbi.nlm.nih.gov/
sites/entrez?db=gene&cmd=
Link&LinkName=gene_pubm
ed&from_uid=227525).
National Center for
Biotechnology Information,
U.S. National Library of
Medicine.
5. Moshous D, Callebaut I, de
Chasseval R, Corneo B,
Cavazzana-Calvo M, Le
Deist F, Tezcan I, Sanal O,
Bertrand Y, Philippe N,
Fischer A, de Villartay JP
(May 2001). "Artemis, a novel
DNA double-strand break
repair/V(D)J recombination
protein, is mutated in human
severe combined immune
deficiency". Cell. 105 (2):
177–86. doi:10.1016/S0092-
8674(01)00309-9 (https://doi.
org/10.1016%2FS0092-867
4%2801%2900309-9).
PMID 11336668 (https://pubm
ed.ncbi.nlm.nih.gov/1133666
8).
gy.org/amigo/term/GO:0006302)
• DNA repair (http://amigo.geneontology.org/amigo/ter
m/GO:0006281)
Sources:Amigo (http://amigo.geneontology.org/) / QuickGO (https://
www.ebi.ac.uk/QuickGO/)
Orthologs
Species Human Mouse
Entrez
64421 (https://www.nc 227525 (https://www.ncb
bi.nlm.nih.gov/entrez/q i.nlm.nih.gov/entrez/quer
uery.fcgi?db=gene&c y.fcgi?db=gene&cmd=retr
md=retrieve&dopt=def ieve&dopt=default&list_u
ault&list_uids=64421 ids=227525&rn=1)
&rn=1)
Ensembl
ENSG00000152457 ENSMUSG00000026648
(http://www.ensembl.o (http://www.ensembl.org/
rg/Homo_sapiens/gen Mus_musculus/genevie
eview?gene=ENSG00 w?gene=ENSMUSG000
000152457;db=core) 00026648;db=core)
UniProt
Q96SD1 (https://www. Q8K4J0 (https://www.uni
uniprot.org/uniprot/Q9 prot.org/uniprot/Q8K4J0)
6SD1)
RefSeq NM_001033855 (https:// NM_001110214 (https://w
(mRNA) www.ncbi.nlm.nih.gov/ ww.ncbi.nlm.nih.gov/entre
entrez/viewer.fcgi?val= z/viewer.fcgi?val=NM_001
NM_001033855) 110214)
NM_001033857 (https:// NM_146114 (https://www.n
www.ncbi.nlm.nih.gov/ cbi.nlm.nih.gov/entrez/vie
entrez/viewer.fcgi?val= wer.fcgi?val=NM_146114)
NM_001033857) NM_175683 (https://www.n
NM_001033858 (https:// cbi.nlm.nih.gov/entrez/vie
www.ncbi.nlm.nih.gov/ wer.fcgi?val=NM_175683)
entrez/viewer.fcgi?val= NM_001302674 (https://w
NM_001033858) ww.ncbi.nlm.nih.gov/entre
NM_001289076 (https:// z/viewer.fcgi?val=NM_001
www.ncbi.nlm.nih.gov/ 302674)
entrez/viewer.fcgi?val= NM_001302684 (https://w
NM_001289076) ww.ncbi.nlm.nih.gov/entre
NM_001289077 (https:// z/viewer.fcgi?val=NM_001
www.ncbi.nlm.nih.gov/ 302684)
entrez/viewer.fcgi?val=
NM_001289077)
6. Li L, Drayna D, Hu D,
Hayward A, Gahagan S,
Pabst H, Cowan MJ (Mar
1998). "The gene for severe
combined immunodeficiency
disease in Athabascan-
speaking Native Americans is
located on chromosome 10p"
(https://www.ncbi.nlm.nih.gov/
pmc/articles/PMC1376812).
Am. J. Hum. Genet. 62 (1):
136–44. doi:10.1086/301688
(https://doi.org/10.1086%2F3
01688). PMC 1376812 (http
s://www.ncbi.nlm.nih.gov/pm
c/articles/PMC1376812).
PMID 9443881 (https://pubme
d.ncbi.nlm.nih.gov/9443881).
7. "Protein Knowledgebase:
Gene DCLRE1C - DNA
cross-link repair 1C protein
(Protein artemis)" (https://ww
w.uniprot.org/uniprot/Q96SD
1). Retrieved June 2, 2011.
8. de Villartay JP (2009). "V(D)J
recombination deficiencies".
Adv. Exp. Med. Biol.
Advances in Experimental
Medicine and Biology. 650:
46–58. doi:10.1007/978-1-
4419-0296-2_4 (https://doi.or
g/10.1007%2F978-1-4419-02
96-2_4). ISBN 978-1-4419-
0295-5. PMID 19731800 (http
s://pubmed.ncbi.nlm.nih.gov/1
9731800).
9. Lu H, Schwarz K, Lieber MR
(2007). "Extent to which
hairpin opening by the
Artemis:DNA-PKcs complex
can contribute to junctional
diversity in V(D)J
recombination" (https://www.n
cbi.nlm.nih.gov/pmc/articles/P
MC2175297). Nucleic Acids
Res. 35 (20): 6917–23.
doi:10.1093/nar/gkm823 (http
s://doi.org/10.1093%2Fnar%2
Fgkm823). PMC 2175297 (htt
ps://www.ncbi.nlm.nih.gov/pm
c/articles/PMC2175297).
PMID 17932067 (https://pubm
ed.ncbi.nlm.nih.gov/1793206
7).
NM_001362903 (https://
NM_001289078 (http www.ncbi.nlm.nih.gov/ent
s://www.ncbi.nlm.nih.g rez/viewer.fcgi?val=NM_
ov/entrez/viewer.fcgi? 001362903)
val=NM_001289078) NM_001362904 (https://
NM_001289079 (http www.ncbi.nlm.nih.gov/ent
s://www.ncbi.nlm.nih.g rez/viewer.fcgi?val=NM_
ov/entrez/viewer.fcgi? 001362904)
val=NM_001289079)
NM_022487 (https://w
ww.ncbi.nlm.nih.gov/e
ntrez/viewer.fcgi?val=
NM_022487)
NM_001350965 (http
s://www.ncbi.nlm.nih.g
ov/entrez/viewer.fcgi?
val=NM_001350965)
NM_001350966 (http
s://www.ncbi.nlm.nih.g
ov/entrez/viewer.fcgi?
val=NM_001350966)
NM_001350967 (http
s://www.ncbi.nlm.nih.g
ov/entrez/viewer.fcgi?
val=NM_001350967)
RefSeq NP_001029027 (https:// NP_001103684 (https://ww
(protein) www.ncbi.nlm.nih.gov/ w.ncbi.nlm.nih.gov/entrez/
entrez/viewer.fcgi?val= viewer.fcgi?val=NP_00110
NP_001029027) 3684)
NP_001029029 (https:// NP_001289603 (https://ww
www.ncbi.nlm.nih.gov/ w.ncbi.nlm.nih.gov/entrez/
entrez/viewer.fcgi?val= viewer.fcgi?val=NP_00128
NP_001029029) 9603)
NP_001029030 (https:// NP_001289613 (https://ww
www.ncbi.nlm.nih.gov/ w.ncbi.nlm.nih.gov/entrez/
entrez/viewer.fcgi?val= viewer.fcgi?val=NP_00128
NP_001029030) 9613)
NP_001276005 (https:// NP_666226 (https://www.n
www.ncbi.nlm.nih.gov/ cbi.nlm.nih.gov/entrez/vie
entrez/viewer.fcgi?val= wer.fcgi?val=NP_666226)
NP_001276005) NP_783614 (https://www.n
NP_001276006 (https:// cbi.nlm.nih.gov/entrez/vie
www.ncbi.nlm.nih.gov/ wer.fcgi?val=NP_783614)
entrez/viewer.fcgi?val=
NP_001276006) NP_001349832 (https://w
ww.ncbi.nlm.nih.gov/entr
ez/viewer.fcgi?val=NP_0
NP_001276007 (http
s://www.ncbi.nlm.nih.g 01349832)
NP_001349833 (https://w
ov/entrez/viewer.fcgi?
10. Povirk LF, Zhou T, Zhou R,
Cowan MJ, Yannone SM
(February 2007). "Processing
of 3'-phosphoglycolate-
terminated DNA double
strand breaks by Artemis
nuclease" (https://doi.org/10.1
074/jbc.M607745200). J. Biol.
Chem. 282 (6): 3547–58.
doi:10.1074/jbc.M607745200
(https://doi.org/10.1074%2Fjb
c.M607745200).
PMID 17121861 (https://pubm
ed.ncbi.nlm.nih.gov/1712186
1).
11. Deckbar D, Birraux J,
Krempler A, Tchouandong L,
Beucher A, Walker S, Stiff T,
Jeggo P, Löbrich M (March
2007). "Chromosome
breakage after G2 checkpoint
release" (http://sro.sussex.ac.
uk/1851/1/deckbar_2007.pdf)
(PDF). J. Cell Biol. 176 (6):
749–55.
doi:10.1083/jcb.200612047
(https://doi.org/10.1083%2Fjc
b.200612047). PMC 2064048
(https://www.ncbi.nlm.nih.gov/
pmc/articles/PMC2064048).
PMID 17353355 (https://pubm
ed.ncbi.nlm.nih.gov/1735335
5).
12. Riballo E, Kühne M, Rief N,
val=NP_001276007) ww.ncbi.nlm.nih.gov/entr
NP_001276008 (http ez/viewer.fcgi?val=NP_0
s://www.ncbi.nlm.nih.g 01349833)
ov/entrez/viewer.fcgi?
val=NP_001276008)
NP_071932 (https://w
ww.ncbi.nlm.nih.gov/e
ntrez/viewer.fcgi?val=
NP_071932)
NP_001337894 (http
s://www.ncbi.nlm.nih.g
ov/entrez/viewer.fcgi?
val=NP_001337894)
NP_001337895 (http
s://www.ncbi.nlm.nih.g
ov/entrez/viewer.fcgi?
val=NP_001337895)
NP_001337896 (http
s://www.ncbi.nlm.nih.g
ov/entrez/viewer.fcgi?
val=NP_001337896)
Location Chr 10: 14.9 – 14.95 Mb Chr 2: 3.42 – 3.46 Mb (http
(UCSC) (https://genome.ucsc.edu/ s://genome.ucsc.edu/cgi-bin/
cgi-bin/hgTracks?org=Hu hgTracks?org=Mouse&db=m
man&db=hg38&position=c m0&position=chr2:3424131-3
hr10:14897359-1495443 464130)
2)
[3] [4]
PubMed
search
Wikidata

Doherty A, Smith GC, Recio

View/Edit Human View/Edit Mouse
MJ, Reis C, Dahm K, Fricke
A, Krempler A, Parker AR,
Jackson SP, Gennery A,
Jeggo PA, Löbrich M
(December 2004). "A
pathway of double-strand
break rejoining dependent
upon ATM, Artemis, and
proteins locating to gamma-
H2AX foci". Mol. Cell. 16 (5):
715–24.
doi:10.1016/j.molcel.2004.10.
029 (https://doi.org/10.1016%
2Fj.molcel.2004.10.029).
PMID 15574327 (https://pubm
ed.ncbi.nlm.nih.gov/1557432
7).
13. Goodarzi AA, Noon AT,
Deckbar D, Ziv Y, Shiloh Y,
Löbrich M, Jeggo PA (July
2008). "ATM signaling
facilitates repair of DNA
double-strand breaks
associated with
heterochromatin". Mol. Cell.
31 (2): 167–77.
doi:10.1016/j.molcel.2008.05.
017 (https://doi.org/10.1016%
2Fj.molcel.2008.05.017).
PMID 18657500 (https://pubm
ed.ncbi.nlm.nih.gov/1865750
0).
14. Beucher A, Birraux J,
Tchouandong L, Barton O,
Shibata A, Conrad S,
Goodarzi AA, Krempler A,
Jeggo PA, Löbrich M
(November 2009). "ATM and
Artemis promote homologous
recombination of radiation-
induced DNA double-strand
breaks in G2" (https://www.nc
bi.nlm.nih.gov/pmc/articles/P
MC2752027). EMBO J. 28
(21): 3413–27.
doi:10.1038/emboj.2009.276
(https://doi.org/10.1038%2Fe
mboj.2009.276).
PMC 2752027 (https://www.n
cbi.nlm.nih.gov/pmc/articles/P
MC2752027).
PMID 19779458 (https://pubm
ed.ncbi.nlm.nih.gov/1977945
8).
15. Woodbine L, Brunton H,
Goodarzi AA, Shibata A,
Jeggo PA (2011).
"Endogenously induced DNA
double strand breaks arise in
heterochromatic DNA regions
and require ataxia
telangiectasia mutated and
Artemis for their repair" (http
s://www.ncbi.nlm.nih.gov/pm
c/articles/PMC3167608).
Nucleic Acids Res. 39 (16):
6986–97.
doi:10.1093/nar/gkr331 (http
s://doi.org/10.1093%2Fnar%2
Fgkr331). PMC 3167608 (http
s://www.ncbi.nlm.nih.gov/pm
c/articles/PMC3167608).
PMID 21596788 (https://pubm
ed.ncbi.nlm.nih.gov/2159678
8).
16. "Entrez Gene: DCLRE1C
DNA cross-link repair 1C
(PSO2 homolog, S.
cerevisiae)" (https://www.ncb
i.nlm.nih.gov/sites/entrez?Db
=gene&Cmd=ShowDetailVie
w&TermToSearch=64421).
17. Ma Y, Pannicke U, Schwarz
K, Lieber MR (March 2002).
"Hairpin opening and
overhang processing by an
Artemis/DNA-dependent
protein kinase complex in
nonhomologous end joining
and V(D)J recombination".
Cell. 108 (6): 781–94.
doi:10.1016/S0092-
8674(02)00671-2 (https://doi.
org/10.1016%2FS0092-867
4%2802%2900671-2).
PMID 11955432 (https://pubm
ed.ncbi.nlm.nih.gov/1195543
2).

Further reading
Dudásová Z, Chovanec M (2003). "Artemis, a novel guardian of the genome". Neoplasma. 50
(5): 311–8. PMID 14628082 (https://pubmed.ncbi.nlm.nih.gov/14628082).
Adams MD, Kerlavage AR, Fleischmann RD, Fuldner RA, Bult CJ, Lee NH, Kirkness EF,
Weinstock KG, Gocayne JD, White O (1995). "Initial assessment of human gene diversity and
expression patterns based upon 83 million nucleotides of cDNA sequence" (http://www.columb
ia.edu/itc/biology/pollack/w4065/client_edit/readings/nature377_3.pdf) (PDF). Nature. 377
(6547 Suppl): 3–174. PMID 7566098 (https://pubmed.ncbi.nlm.nih.gov/7566098).
Wood RD, Mitchell M, Sgouros J, Lindahl T (2001). "Human DNA repair genes" (https://doi.org/
10.1126/science.1056154). Science. 291 (5507): 1284–9. doi:10.1126/science.1056154 (http
s://doi.org/10.1126%2Fscience.1056154). PMID 11181991 (https://pubmed.ncbi.nlm.nih.gov/11
181991).
Ma Y, Pannicke U, Schwarz K, Lieber MR (2002). "Hairpin opening and overhang processing
by an Artemis/DNA-dependent protein kinase complex in nonhomologous end joining and
V(D)J recombination". Cell. 108 (6): 781–94. doi:10.1016/S0092-8674(02)00671-2 (https://doi.o
rg/10.1016%2FS0092-8674%2802%2900671-2). PMID 11955432 (https://pubmed.ncbi.nlm.ni
h.gov/11955432).
Li L, Moshous D, Zhou Y, Wang J, Xie G, Salido E, Hu D, de Villartay JP, Cowan MJ (2002). "A
founder mutation in Artemis, an SNM1-like protein, causes SCID in Athabascan-speaking
Native Americans" (https://doi.org/10.4049/jimmunol.168.12.6323). J. Immunol. 168 (12): 6323–
9. doi:10.4049/jimmunol.168.12.6323 (https://doi.org/10.4049%2Fjimmunol.168.12.6323).
PMID 12055248 (https://pubmed.ncbi.nlm.nih.gov/12055248).
Callebaut I, Moshous D, Mornon JP, de Villartay JP (2002). "Metallo-beta-lactamase fold within
nucleic acids processing enzymes: the beta-CASP family" (https://www.ncbi.nlm.nih.gov/pmc/a
rticles/PMC134238). Nucleic Acids Res. 30 (16): 3592–601. doi:10.1093/nar/gkf470 (https://doi.
org/10.1093%2Fnar%2Fgkf470). PMC 134238 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC
134238). PMID 12177301 (https://pubmed.ncbi.nlm.nih.gov/12177301).
Noordzij JG, Verkaik NS, van der Burg M, van Veelen LR, de Bruin-Versteeg S, Wiegant W,
Vossen JM, Weemaes CM, de Groot R, Zdzienicka MZ, van Gent DC, van Dongen JJ (2003).
"Radiosensitive SCID patients with Artemis gene mutations show a complete B-cell
differentiation arrest at the pre-B-cell receptor checkpoint in bone marrow" (http://repub.eur.nl/p
ub/8235). Blood. 101 (4): 1446–52. doi:10.1182/blood-2002-01-0187 (https://doi.org/10.1182%
2Fblood-2002-01-0187). PMID 12406895 (https://pubmed.ncbi.nlm.nih.gov/12406895).
Moshous D, Pannetier C, de Chasseval R, le Deist F, Cavazzana-Calvo M, Romana S,
Macintyre E, Canioni D, Brousse N, Fischer A, Casanova JL, de Villartay JP (2003). "Partial T
and B lymphocyte immunodeficiency and predisposition to lymphoma in patients with
hypomorphic mutations in Artemis" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC151863). J.
Clin. Invest. 111 (3): 381–7. doi:10.1172/JCI16774 (https://doi.org/10.1172%2FJCI16774).
PMC 151863 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC151863). PMID 12569164 (https://
pubmed.ncbi.nlm.nih.gov/12569164).
Kobayashi N, Agematsu K, Sugita K, Sako M, Nonoyama S, Yachie A, Kumaki S, Tsuchiya S,
Ochs HD, Sugita K, Fukushima Y, Komiyama A (2003). "Novel Artemis gene mutations of
radiosensitive severe combined immunodeficiency in Japanese families". Hum. Genet. 112 (4):
348–52. doi:10.1007/s00439-002-0897-x (https://doi.org/10.1007%2Fs00439-002-0897-x).
PMID 12592555 (https://pubmed.ncbi.nlm.nih.gov/12592555).
Kobayashi N, Agematsu K, Nagumo H, Yasui K, Katsuyama Y, Yoshizawa K, Ota M, Yachie A,
Komiyama A (2003). "Expansion of clonotype-restricted HLA-identical maternal CD4+ T cells
in a patient with severe combined immunodeficiency and a homozygous mutation in the
Artemis gene". Clin. Immunol. 108 (2): 159–66. doi:10.1016/S1521-6616(03)00095-0 (https://do
i.org/10.1016%2FS1521-6616%2803%2900095-0). PMID 12921762 (https://pubmed.ncbi.nlm.
nih.gov/12921762).
Poinsignon C, Moshous D, Callebaut I, de Chasseval R, Villey I, de Villartay JP (2004). "The
metallo-beta-lactamase/beta-CASP domain of Artemis constitutes the catalytic core for V(D)J
recombination" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2211804). J. Exp. Med. 199 (3):
315–21. doi:10.1084/jem.20031142 (https://doi.org/10.1084%2Fjem.20031142). PMC 2211804
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2211804). PMID 14744996 (https://pubmed.ncb
i.nlm.nih.gov/14744996).
 Pannicke U, Ma Y, Hopfner KP, Niewolik D, Lieber MR, Schwarz K (2004). "Functional and
biochemical dissection of the structure-specific nuclease ARTEMIS" (https://www.ncbi.nlm.nih.
gov/pmc/articles/PMC404326). EMBO J. 23 (9): 1987–97. doi:10.1038/sj.emboj.7600206 (http
s://doi.org/10.1038%2Fsj.emboj.7600206). PMC 404326 (https://www.ncbi.nlm.nih.gov/pmc/arti
cles/PMC404326). PMID 15071507 (https://pubmed.ncbi.nlm.nih.gov/15071507).
Zhang X, Succi J, Feng Z, Prithivirajsingh S, Story MD, Legerski RJ (2004). "Artemis is a
phosphorylation target of ATM and ATR and is involved in the G2/M DNA damage checkpoint
response" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC517881). Mol. Cell. Biol. 24 (20):
9207–20. doi:10.1128/MCB.24.20.9207-9220.2004 (https://doi.org/10.1128%2FMCB.24.20.920
7-9220.2004). PMC 517881 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC517881).
PMID 15456891 (https://pubmed.ncbi.nlm.nih.gov/15456891).
Poinsignon C, de Chasseval R, Soubeyrand S, Moshous D, Fischer A, Haché RJ, de Villartay
JP (2004). "Phosphorylation of Artemis following irradiation-induced DNA damage". Eur. J.
Immunol. 34 (11): 3146–55. doi:10.1002/eji.200425455 (https://doi.org/10.1002%2Feji.2004254
55). PMID 15468306 (https://pubmed.ncbi.nlm.nih.gov/15468306).

External links
Overview of all the structural information available in the PDB for UniProt: Q96SD1 (https://ww
w.ebi.ac.uk/pdbe/pdbe-kb/proteins/Q96SD1) (Protein artemis) at the PDBe-KB.

Retrieved from "https://en.wikipedia.org/w/index.php?title=Artemis_(protein)&oldid=950372207"

This page was last edited on 11 April 2020, at 18:58 (UTC).

Text is available under the Creative Commons Attribution-ShareAlike License; additional terms may apply. By using this
site, you agree to the Terms of Use and Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia
Foundation, Inc., a non-profit organization.